Forty years of study on interactions between walnut tree and arbuscular mycorrhizal fungi. A review

AbstractWalnut trees are among the most important hardwood species in the northern hemisphere, ecologically and economically. They are mainly cultivated for timber and nut production but are also attractive ornamental trees in parks. Establishing walnut orchards is difficult because seedlings have a coarse root architecture and few of them survive to transplanting. Planting success is mainly determined by the root system morphology and the nutrient status of the seedlings, so that rhizosphere conditions are critical for plant performance. Walnut trees can associate with soil-borne arbuscular mycorrhizal fungi, which are obligate biotrophs. In this association, plant-produced carbon compounds are traded against fungus-acquired soil mineral nutrients. The beneficial effect of arbuscular mycorrhizal symbiosis on hardwood seedling quality and field performance has long been known, but an integrated view is lacking about the effects of arbuscular mycorrhizas on walnut cropping. Therefore, we surveyed the literature published over the last 40 years to provide up-to-date knowledge on the relationships between arbuscular mycorrhizas and walnut trees. Our review outlines the major following points: (1) the arbuscular-mycorrhiza-mediated nutrient uptake capacity of walnut trees is associated with first- to third-order roots, and fibrous tip-ended roots are dependent on arbuscular mycorrhizal fungi, whereas pioneer roots are not; (2) early inoculation with arbuscular mycorrhizal fungi improves the survival and seedling performance attributes of transplanted walnut trees: biotization enhances walnut transplant success by increasing the number of lateral roots and plant P uptake, but these benefits are fungus- and host-dependent; (3) in the context of walnut agroforestry, deeply rooted walnut trees play a role as reservoirs of arbuscular mycorrhizal fungal propagules for the surrounding vegetation, but tree shade and soluble phosphate availability decrease walnut mycorrhizal dependency; and (4) the arbuscular mycorrhizal mycelium mediates the transport of juglone and thus plays a role in walnut tree allelopathy.